Isolated dinosaur teeth recovered from seven localities near Fuxin (western Liaoning Province, northeastern China) are described. They come from sediments belonging to the Shahai and Fuxin formations, considered to be Aptian to Albian in age. Seven taxa have been recognized. They include the oviraptorosaur Incisivosaurus, dromaeosaurid theropods, Euhelopus-like sauropods, as well as indeterminate nodosaurid, ankylosaurid, iguanodontoid and basal neoceratopsian ornithischians. The Shahai and Fuxin dinosaur faunas show the persistence of some Jehol biota taxa such as the highly specialised Incisivosaurus, basal titanosauriform sauropods, basal neoceratopsians and some dromaeosaurids, and the addition of more derived iguanodontoids and ankylosaurians. The persistence of some dinosaurs of the Jehol Biota into the Shahai and Fuxin formations suggests a long term stability of Liaoning terrestrial environments during the Early Cretaceous. Despite sampling bias and the rather small sample that must be taken into account, teeth abundances show a significant compositional difference between the localities of the Shahai and Fuxin formations, neoceratopsian teeth representing one third of dinosaur tooth remains in the Shahai Formation whereas they are totally absent in the Fuxin Formation. Ankylosaur teeth, in contrast, represent 3 percent of total remains in the Shahai Formation, whereas they seem to be the only herbivorous dinosaurs in the Fuxin Formation with 40 percent of the total number of teeth (the rest being theropod dinosaur teeth). Although a difference in micro-environmental conditions between Shahai and Fuxin localities may at least partly explain such pattern, the unusual and unbalanced faunal composition of Fuxin localities remains enigmatic and will need further field collecting in order to be clarified.

Terrestrial reptile remains are very rare in the Lower Cretaceous of South Australia, but include the holotype of the small theropod Kakuru. Here, we review this taxon and other archosaur specimens collected from the Bulldog Shale (Aptian) of Andamooka and Coober Pedy. Kakuru possesses no unique characters or character state combinations and is regarded as a nomen dubium, representing an indeterminate tetanuran theropod. Two other specimens (a left metatarsal and astragalus) can be referred to Dinosauria, but the identity of several other specimens (phalanges and a centrum) can only be resolved to the level of an indeterminate archosaur.

Since 2002 a number of sites containing stegosaurian remains (bones and tracks) have been discovered in the Villar del Arzobispo Formation (Tithonian–Berriasian) in the Province of Teruel, Spain, mostly in the areas of El Castellar and Riodeva. The material from the latter consists of the postcranial remains of the axial, pelvic and appendicular skeleton of several different sized specimens related to the genus Dacentrurus. The footprints made by a stegosaurian at El Castellar site reflect a new type of medium gauge trackway: Deltapodus ibericus isp. nov. The presence of these tracks near abundant bones related to Dacentrurus highlights this geological formation as a window through which to examine the systematics, behaviour and palaeoecology of these thyreophoran dinosaurs. Since these fossil bones were discovered next to those of other sauropods and ornithopods (Turiasaurus and Ornithopoda indet.) where there has been virtually no transport of the remains, and since stegosaurian tracks have been found in ichnoassociations with the same groups, they may have coexisted in the wetlands of restricted tidal environments during the Tithonian–Berriasian. The presence of common (or at least phylogenetically closely related) taxa and ichnotaxa in Western Europe, North America and Africa indicates that sea–level fall episodes may have occurred during which the fauna of each area may have reached the emerged regions of the others during the Late Jurassic.

The holotype of Brachylophosaurus canadensis was collected by C.M. Sternberg in 1936 from the Oldman Formation (Upper Cretaceous) of Alberta, Canada, with a brief description focusing on the cranium following in 1953. A re-description of the holotype skeleton confirms its position as a hadrosaurine dinosaur exhibiting the following unique combination of characters: paddle-like solid nasal crest; posteriorly elongated prefrontal; only the anterior tip of the lacrimal contacting maxilla; extremely elongated anterior maxillary process; and a quadratojugal with a noncrescentic posterior margin variably forming a paraquadratic foramen with the quadrate. Comparison of the holotype with other specimens assigned to the taxon reveals numerous morphological differences. These variable features include the presence/absence of an accessory premaxillary foramen, the number of maxillary/dentary vertical tooth rows, the number of functional teeth per tooth row contributing to the dental occlusal surface, the bones forming the mandibular glenoid, the presence/absence of a nasal foramen, nasal crest morphology, the number of cervical vertebrae, and axis morphology.

The cranial anatomy of the helmet-crested lambeosaurine Hypacrosaurus altispinus (Ornithischia: Hadrosauridae) is described, with a focus on ontogenetic and individual variation in phylogenetically significant characters of the cranial crest, braincase, and facial skeleton. Cranial material of H. altispinus represents a relatively complete growth series that includes crestless juveniles of less than half the size of large individuals with fully developed crests. Cranial ontogeny is compared with other lambeosaurines using bivariate morphometrics and through qualitative comparison of a size-standardized cranial growth series. Bivariate analyses reveal that the relative growth of the skull and cranial crest of H. altispinus and H. stebingeri are similar, and that Hypacrosaurus more closely resembles Corythosaurus than Lambeosaurus. Hypacrosaurus altispinus is systematically revised. The taxon is characterized by five autapomorphies, most of which are concentrated in the skull, as well as an enlarged terminal ischial foot. Maximum parsimony and Bayesian likelihood (Mk+gamma) phylogenetic analyses were conducted to test the monophyly of the genus. Hypacrosaurus monophyly is corroborated in light of new anatomical data. Although H. stebingeri and H. altispinus share few derived characters of the skull, the hypothesis that H. stebingeri is a metaspecies that represents the ancestor of H. altispinus cannot be rejected.

Pterosaur remains are very rare in Australasia and especially in Upper Cretaceous strata. Thus, the discovery of a jaw fragment from the Cenomanian-Coniacian Molecap Greensand near Gingin in Western Australia represents an important new stratigraphical occurrence for the region. Although the teeth are not preserved, the presence of labio-lingually compressed alveoli that are anterolaterally oriented, variable in shape/size (inferring heterodonty) and very widely spaced is reminiscent of ornithocheirids—a geographically cosmopolitan clade of predominantly Early Cretaceous pterodactyloids. If correct, this identification could extend the known range of Ornithocheiridae through to the Late Cretaceous in the Southern Hemisphere.

New isolated pterodactyloid bones from the Toolebuc Formation are described. The first one consists of a complete wing metacarpal 212 mm long, representing an individual with an estimated wing span of 4 m. Small depressions on the anterior surface are present and represent tooth marks showing that this specimen was subjected to scavenging prior to fossilization. The other bone consists of a three-dimensionally preserved cervical vertebra lacking most of the neural arch. The specimens are clearly referable to the derived pterosaur clade Pterodactyloidea. Based on several features such as the position of the pneumatic foramen and the particular shape and proportions of those elements, they possibly are members of, or closely related to, the Anhangueridae. The record of the Australian pterosaurs is reviewed here and represents the known southern distributional limit for Cretaceous pterosaurs, arguing against some older ideas of a more geographically restricted range for these flying reptiles.

Three specimens of sauropod dinosaur skin impressions from footprints in the Lower Cretaceous Jindong and Haman formations are unusual because they are not associated with well-preserved ‘host’ tracks. This suggests a hitherto unreported mode of preservation in which diagnostic but isolated skin traces may appear associated with substrates where only the superficial layers were originally soft. Two specimens from the Haman Formation of Gainri and Sinsu Island reveal large pentagonal to heptagonal scale impressions (size 2.0 – 2.5 cm). Another specimen from the Jindong Formation of the Deokmyeongri area is an extremely well-preserved, honeycomb-like pattern of hexagonal scale impressions, but an adjacent part reveals pentagonal to heptagonal polygons of variable size. Collectively these skin impressions more than double the sparse record of sauropod skin impressions and show that patterns (morphology) are consistent from the Jurassic to the Cretaceous. Although comprehensive syntheses have yet to be attempted, skin traces are amendable to systematic description using tubercle size, shape, thickness, and inter-tubercle groove thickness. Such features are useful for distinguishing the different morphologies of dinosaurs and other vertebrates that lived in Korea and other regions during the Mesozoic.

Dinosaur skin impressions have occasionally been confused with glypogryptids, typical deep sea feeding traces, small desiccation cracks, and small load casts which they superficially resemble. To avoid such confusion we enumerate criteria useful for distinguishing dinosaur skin impressions from similar shaped invertebrate trace fossils and inorganic sedimentary structures. These criteria vary in importance but include preservation, morphology, dimension of morphology, depositional environment, deformation of underlying sediments, uniformity of size and shape of individual scale impressions within single specimens, ornamentation, and geological age.

We present the first detailed description of Perudyptes devriesi, a basal penguin from the middle Eocene (~42 Ma) Paracas Formation of Peru, and a new analysis of all published extinct penguin species as well as controversial fragmentary specimens. The Perudyptes devriesi holotype includes key regions of the skull and significant postcranial material, thus helping to fill a major phylogenetic and stratigraphic (~20 million year) gap between the earliest fossil penguins (Waimanu manneringi and Waimanu tuatahi, ~58–61.6 Ma) and the next oldest partial skeletons. Perudyptes devriesi is diagnosable by five autapomorphies: (1) an anteroventrally directed postorbital process, (2) marked anterior expansion of the parasphenoid rostrum, (3) posterior trochlear ridge of the humerus projecting distal to the middle trochlear ridge and conformed as a large, broadly curved surface, (4) convex articular surface for the antitrochanter of the femur, and (5) extremely weak anterior projection of the lateral condyle of the tibiotarsus. The skull of Perudyptes is characterized by deep temporal fossae and an elongate, narrow beak that differs from other reported stem penguins in its short mandibular symphysis. The wing skeleton of Perudyptes preserves a combination of plesiomorphic features also observed in the basal penguin Waimanu and derived features shared with more crownward penguins. Features of the wing optimized as primitive for Sphenisciformes include retention of a discrete dorsal supracondylar tubercle on the humerus and presence of a modestly projected pisiform process on the carpometacarpus. Derived features present in Perudyptes and all more crownward penguins, but absent in Waimanu, include a more flattened humerus, development of a trochlea for the tendon of m. scapulotriceps at the distal end of the humerus, and bowing of the anterior face of the carpometacarpus.
A combined molecular and morphological dataset for Spheniciformes was expanded by adding 25 osteological and soft tissue characters as well as 11 taxa. In agreement with previous results, Perudyptes devriesi is identified as one of the most basal members of Sphenisciformes. This analysis also confirms the placement of the middle/late Miocene (~11–13 Ma) fossil Spheniscus muizoni as a member of the Spheniscus clade and places the late Miocene (~10 Ma) Madrynornis mirandus as sister taxon to extant Eudyptes. These two species, known from relatively complete partial skeletons, are the oldest crown clade penguin fossils and represent well-corroborated temporal calibration points for the Spheniscus-Eudyptula divergence and Megadyptes-Eudyptes divergence, respectively. Our results reaffirm that the Miocene penguin taxon Palaeospheniscus, recently proposed to represent a member of the crown radiation, belongs outside of the crown clade Spheniscidae.
The phylogenetic positions of small Eocene Antarctic penguin taxa (Delphinornis, Marambiornis, and Mesetaornis) recently proposed as possible direct ancestors to crown Spheniscidae were further evaluated using alternate coding strategies for incorporating scorings from isolated elements that preserve critical morphologies and are thought to represent these taxa, although they cannot yet be reliably assigned to individual species. Under all scoring regimes, Delphinornis, Marambiornis, and Mesetaornis were recovered as distantly related to Spheniscidae.
Using synapomorphies identified in the primary analysis, we evaluated the phylogenetic position of fragmentary specimens, including the holotypes of valid but poorly known species, specimens currently unassignable to the species level, and morphologically distinct specimens that have not yet been named. All pre-Miocene specimens can be excluded from Spheniscidae based on presence of plesiomorphies lost in all crown penguins, consistent with a recent radiation for the penguin crown clade. This study provides additional support for a scenario of penguin evolution characterized by an origin of flightlessness near the K-T boundary, dispersal throughout the Southern Hemisphere during the early Paleogene, and a late Cenozoic origin for the crown clade Spheniscidae. Stratigraphic distribution and phylogenetic relationships of fossil penguins are consistent with distinct radiations during the Eocene, Oligocene, and Miocene. While the Eocene and Oligocene penguin faunas are similar in many respects, the Miocene fauna is characterized by smaller average size and novel cranial morphologies, suggesting that an ecological shift in diet occurred close to the origin of crown Spheniscidae.

The feeding apparatus of Paleogene birds is rarely well-preserved. Here, we describe the earliest known pelican (early Oligocene, Luberon, southeastern France), with its almost complete beak. Morphologically identical to modern pelicans, the new fossil already shows several advanced features unique to extant species of the genus Pelecanus. It probably belongs to the lineage ancestral to all or some of these pelican species. This fossil reveals a remarkable evolutionary stasis in the morphology of such an advanced avian feeding apparatus through ca. 30 million years. Several hypotheses are proposed to suggest explanations for such examples of long stases in volant homeothermic vertebrates.

The Cameros Basin sedimentary infill comprises a large, essentially continental, megasequence ranging from the Tithonian (Upper Jurassic) to the Albian (Lower Cretaceous). It occupies an area of some 5500 km2, and is home to around 300 dinosaur tracksites. Most of these tracksites are found in either the Huérteles Formation, which is part of the Oncala Group (Berriasian), or the Enciso Group (Lower Aptian), both of which represent early Cretaceous lacustrine episodes. Dinosaur trackways (n=1170) from both episodes were analysed in order to establish the preferred direction of dinosaur movement, and to determine whether these movements were influenced by the paleogeographic and palaeoenvironmental conditions of the area.

The Huérteles Formation is interpreted as a complex record of alluvial plain systems distally connected with a playa-lake. Its dinosaur tracksites are distributed throughout its alluvial plain facies and trackways show two preferential unidirectional orientations: 1) NW, more or less parallel to the distribution of the facies belt and 2) NNE. The Enciso Group is represented by a wide and shallow lacustrine system connected with marine environments towards the SE (Iberian Basin realm) and in close proximity to marine settings to the NW (Basque-Cantabrian Basin realm), rendering the Cameros Basin as the only continental connection between the Ebro and Iberian Massifs. The fluctuating (but always shallow) water level of the system with frequent desiccations probably allowed dinosaurs to pass through the lake basin. As a consequence of these paleogeographic restrictions, the dinosaur trackways generally show a bidirectional NE—SW orientation. Despite the temporal and geographical differences between the Huérteles Formation and Enciso Group, the ichnocenoses of both are dominated by theropod dinosaur trackways (85% as a mean value). This is probably explained by these dinosaurs being more active than others, a consequence of their searching/hunting behaviour.

'Bizarre structures' in dinosaurs have four main traditional explanations: mechanical function, sexual selection, social selection and species recognition. Any of these can be plausible for individual species, but they fail to be persuasive when other lines of evidence cannot adequately test them. The first three also fail as general propositions when phylogenetic analyses based on other characters do not support scenarios of selective improvement of such functions in their clade (or the explanation simply does not apply to any other species in the clade). Moreover, the hypothesis of sexual selection requires significant sexual dimorphism, which has never been conclusively established in dinosaurs.

We propose instead that species recognition may have been a more general force that drove the evolution of bizarre structures in dinosaurs. That is, the bizarre structures communicate to other individuals a variety of possible associational cues, including species identification, potential protection and social habits and the appropriateness of potential mates. In other words, bizarre structures amount to an advertisement for positive association. Neither species recognition nor any other hypothesis should be a 'default' explanation. Although direct observation is impossible, we propose two tests. First, contrary to adaptive, social or sexual selection, under the species recognition model morphology should be expected to evolve without obvious directional trends, because the only objective is to differ from one's relatives. Hence, patterns of evolution of bizarre structures should be relatively proliferative and non-directional. Second, several contemporaneous species should overlap in geographic range (sympatric, parapatric, peripatric). Fossil species often show evidence of this pattern in the past by 'ghost ranges' of related taxa. These tests together could reinforce or weaken an argument for species recognition.

Hadrosaurids were the most derived ornithopods and amongst the most diverse herbivore dinosaurs during the Late Cretaceous of Europe, Asia, and the two Americas. Here, their biogeographical history is reconstructed using dispersal-vicariance analysis (DIVA). The results showed that Hadrosauridae originated in North America and soon after dispersed to Asia no later than the Late Santonian. The most recent common ancestor of Saurolophidae (= Saurolophinae + Lambeosaurinae) is inferred to have been widespread in North America and Asia. The split between saurolophines and lambeosaurines occurred in response to vicariance no later than the Late Santonian: the former clade originated in North America, whereas the latter did so in Asia. Saurolophine biogeographical history included a minimum of five dispersal events followed by vicariance. Four of these dispersals were inferred to have occurred from North America to Asia during the Campanian and Early Maastrichtian, whereas a fifth event represented a southward dispersal from North to South America no later than the Late Campanian. The historical biogeography of lambeosaurines was characterized by an early evolution in Asia, with a Campanian dispersal to the European archipelago followed by vicariance. Reconstruction of the ancestral areas for the deepest nodes uniting the more derived lambeosaurines clades ('hypacrosaurs', 'corythosaurs', and 'parasaurolophs') is ambiguous. The split between North American and Asian clades of 'hypacrosaurs' and 'parasaurolophs' occurred in response to vicariance during the Campanian. The evolutionary history of North American 'hypacrosaurs' and 'parasaurolophs' was characterized by duplication events. The latter also characterized the Late Campanian 'corythosaurs', which remained restricted to North America.

The late Cretaceous hadrosaurids were the most specialized and diverse clade of ornithopod dinosaurs. Parsimony and Bayesian methods were implemented to elucidate the phylogenetic relationships of all hadrosaurid species. Traditional and geometric morphometrics were applied to discover patterns of variation containing phylogenetic information. In total, 286 phylogenetically informative characters (196 cranial and 90 postcranial) were defined and documented: the most extensive character data set ever constructed for hadrosaurid dinosaurs. Of these, 136 characters were used for the first time in phylogenetic analysis of these ornithopods, and 93 were modified from those of other authors. Parsimony and the Bayesian analysis (using the Mk model without the gamma parameter) confirmed the split of hadrosaurids into Saurolophinae and Lambeosaurinae. Saurolophines included a major clade composed of the Prosaurolophus–Saurolophus and the Kritosaurus–Gryposaurus–Secernosaurus subclades. Edmontosaurus and Shantungosaurus were recovered outside the major clade of saurolophines. The Brachylophosaurus clade was recovered as the most basal clade of saurolophines in the parsimony analysis, whereas following the Bayesian analysis it was recovered as the sister clade to the Kritosaurus–Gryposaurus–Secernosaurus clade. These two analyses resulted in a Lambeosaurinae composed of a succession of Eurasian sister taxa to two major clades: the Parasaurolophus clade and the Hypacrosaurs altispinus–Corythosaurus clade. In contrast, the Bayesian analysis using the Mk model with the gamma parameter included, resulted in an unbalanced hadrosauroid tree, with a paraphyletic Saurolophinae, and with the Prosaurolophus clade, Edmontosaurus, and Shantungosaurus as successively closer sister taxa to Lambeosaurinae. Based on the strict reduced consensus tree derived from the parsimony analysis, Hadrosauridae was redefined as the clade stemming from the most recent common ancestor of Hadrosaurus foulkii and Parasaurolophus walkeri.

Finely-bedded lacustrine deposits of the Lower Cretaceous (Aptian) Xiagou Formation exposed in the Changma Basin of Gansu Province, northwestern China, have yielded numerous fossil vertebrate remains, including approximately 100 avian specimens. Though the majority of these birds appear referable to the ornithuromorph Gansus yumenensis, a number of enantiornithine fossils have also been recovered. Here we report on a specimen consisting of a complete, three-dimensionally preserved sternum, furcula, and sternal ribs that represents a second ornithuromorph taxon from the Xiagou Formation at Changma. The fossil exhibits morphologies that distinguish it from all previously-known Xiagou birds and demonstrate that it represents a derived non-ornithurine member of Ornithuromorpha. Though it is morphologically distinct from the equivalent elements of all other described ornithuromorphs, the material is too incomplete to justify the erection of a new taxon. Nonetheless, it increases the taxonomic diversity of the Xiagou avifauna, thereby expanding our knowledge of Early Cretaceous avian diversity and evolution.
Zhou, C.-F., Gao, K.-Q. & Fox, R. C. 2010. Morphology and histology of lattice-like ossified epaxial tendons in Psittacosaurus (Dinosauria: Ceratopsia). Acta Geologica Sinica (English Edition) 84(3): 463-471.

Epaxial tendons play an important role in the study of the musculoskeletal system and locomotory style of dinosaurs. Although the ossified epaxial tendon lattice is fairly well known in Iguanodontoidea, only recently has knowledge of this complex been extended to ceratopsians. This study concerns the gross morphology and microstructure of the tendon lattice in Psittacosaurus, a basal ceratopsian. As in the neoceratopsian Chasmosaurus, the ossified tendons of Psittacosaurus form a three-layered, lattice-like structure. The microstructure of the tendons in large psittacosaur individuals retains an early stage of ossification, as in juvenile birds and nestling hadrosaurs, suggesting a slow developmental rate of ossification of the tendons in psittacosaur ontogeny. Comparative study indicates that a lattice-like arrangement of three-layered epaxial tendons is widely distributed in Cerapoda. This pattern also extends to Ankylosauria, implying a similar pattern of the epaxial muscles through the ornithischian clade. In addition, comparison with crocodiles implies that the different morphology of ossified tendons in dinosaurs may be associated with adaptive aspects of their paleobiology, not simply a side effect of skeletal ossification. In contrast to the short tendons in quadrupedal Chasmosaurus and Protoceratops, the elongated tendons in Psittacosaurus may be related to the bipedal locomotion characteristic of this taxon.